Description

Family Fagaceae

Trees
or rarely shrubs
, monoecious, evergreen
or deciduous. Stipules usually early deciduous. Leaves alternate, sometimes false-whorled in Cyclobalanopsis. Inflorescencesunisexual
or androgynous with female cupules at the base
of an otherwise male inflorescence. Male inflorescences a penduloushead
or erect
or pendulous catkin, sometimes branched; flowers in dense cymules
. Male flower: sepals 4-6(-9), scalelike, connate
or distinct
; petals absent; filamentsfiliform
; anthersdorsifixed
or versatile, opening by longitudinal
slits; with or without a rudimentarypistil. Female inflorescences of 1-7 or more flowers subtended individually or collectively by a cupule formed from numerous
fused bracts, arranged individually or in small groups along an axis or at base of an androgynous inflorescence or on a separate axis. Female flower: perianth 1-7 or more; pistil 1; ovaryinferior, 3-6(-9) -loculed; style and carpels as many as locules; placentationaxile
; ovules 2 per locule. Fruit a nut. Seed usually solitary by abortion
(but may be more than 1 in Castanea, Castanopsis, Fagus, and Formanodendron), without endosperm; embryo large.

Seven to 12 genera (depending on interpretation) and 900-1000 species: worldwide except for tropical
and S Africa; seven genera and 294 species (163 endemic, at least three introduced
) in China.

Many species are important timber trees. Nuts of Fagus, Castanea, and of most Castanopsis species are edible, and oil
is extracted from nuts of Fagus. Nuts of most species of this family
contain copious
amounts of water soluble tannin. Members
of the Fagaceae are the main element
of both broad-leaved evergreen and mixed mesophyticforests
from 500-3200 m.[1]

Quercus is without doubt one of the most important woody genera of the Northern Hemisphere. Historically, oaks have been an important source of fuel, fodder
, and building materials
throughout their range
. Other products include tannins and dyes, and oak bark
and leaves were often used for tanning leather
. Acorns were historically an important food for indigenous people in North America, Central America, Europe, and Asia. In some areas, acorn consumption
is still important, but in general, because of the intense preparation necessary to remove tannins and strong
flavor of acorn products, they have fallen out of use as human food in developed areas. They do remain, however, an important mast for wildlife and domesticated animals in many rural areas.

Among the most important diagnostic characters within Quercus, and particularly the white oak group ( Quercus sect. Quercus), are features of the foliartrichomes
. Often these can be seen with a 10× or 15× hand lens
; higher magnifications are sometimes required and are useful particularly when characters for a species or complex
are first studied and mastered for later use in the field
. Although these microscopic characters may seem intimidating, the alternative characters of leaf shape and dentition, so often used in the field, are unreliable in many cases. The large number of misidentified specimens in herbaria that can be easily identified properly with the use of trichome characters illustrates this point
. Additionally, many specimens are encountered, both in field and herbarium
, that lack fruit or have only immature
fruit. Very few species require mature
fruit for proper diagnosis
; most can be adequately identified with a representative selection of mature sun leaves attached, if possible, to twigs
with mature buds. The combination
of leaf vestiture, form of the margin (entire, lobed, toothed, spinose
), twigvestiture
, and bud form and vestiture constitute the majority of diagnostic features minimally required at species level.

Staminate floral and inflorescence characters have not been used to any significant extent in the taxonomy of Quercus . Immature, flowering material is often difficult to identify with certainty, and floral
features such as number and form of sepals, number of stamens, and pubescence
of flowers or floral rachises seem to vary independently of species affinity within many groups. Because of these problems, descriptions
of staminate features are excluded
in this treatment as unreliable and of little diagnostic value. When collecting flowering oaks, make a point of gathering fallen fruit and mature leaves carefully from the ground
, if available, and revisit such populations again when mature material is available to verify identifications.

The character of acorn maturation in the first year (annual maturation) or second year (biennial maturation) after pollination is commonly used to differentiate major groups within Quercus . All of the North American white oaks have annual maturation; all of the Protobalanus group have biennial maturation; and the vast majority of red oaks have biennial maturation, with one eastern North American and a few western species with annual maturation. In the field, this character can be observed throughout the growing season
by examining a sample
of twigs from the same tree. If developing fruits exhibit
a single size class
and are found only on the current
year's growth, maturation is annual; if the developing fruits exhibit two size classes with small pistillate flowers on new growth and larger developing fruit on the previous year's twigs, maturation is biennial. In Quercus sect. Protobalanus, biennial maturation may be mistaken for annual maturation because all of the species are fully evergreen, and often the twigs bearing fruit do not produce
new growth in the second year after pollination. In such cases, careful examination of a broad sample of twigs from within one tree and throughout a population will verify biennial maturation. Herbarium specimens are sometimes inadequate for this determination.

Hybridization among species of Quercus has been widely documented and even more widely suspected. An astounding number of hybrid combinations have been reported in the literature, and many of these have been given species names
, either before or after their hybrid status was known (E. J. Palmer 1948) . Hybrids are known to occur in the wild only between members
of the same section, and attempts at artificial crosses between species from different sections or subgenera
within Quercus have failed with very few exceptions (W. P. Cottam et al.
1982) .

Hybridization in most cases results in solitary unusual trees or scatteredclusters
of intermediate individuals (J. W. Hardin 1975) . In some cases, however, populations of fairly broad distribution and extreme variability, often with a majority of intermediate types, may occur. Such instances occur in both the red oak and white oak groups, and to a lesser extent in the Protobalanus group.

When dealing with a suspected hybrid, therefore, one should first consider the possibility of intraspecificvariation
or environmental plasticity, and then seek parentage among sympatric members of the same section. Because of the almost infinite number of possible hybrid combinations, and the myriad names
applied to them, only those that appear to be prominent
either locally or in widespread areas are dealt with here. The interested reader is referred to various discussions of oak hybridization in the literature (e.g.
, E. J. Palmer 1948; J. W. Hardin 1975) .[2]

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